{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T11:18:12Z","timestamp":1789816692532,"version":"4.0.1"},"reference-count":77,"publisher":"Wiley","issue":"6","license":[{"start":{"date-parts":[[2012,3,30]],"date-time":"2012-03-30T00:00:00Z","timestamp":1333065600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["The Anatomical Record"],"published-print":{"date-parts":[[2012,6]]},"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Narwhal tusks, although well described and characterized within publications, are clouded by contradictory references, which refer to them as both incisors and canines. Vestigial teeth are briefly mentioned in the scientific literature with limited descriptions and no image renderings. This study first examines narwhal maxillary osteoanatomy to determine whether the erupted tusks are best described as incisiform or caniniform teeth. The study also offers evidence to support the evolutionary obsolescence of the vestigial teeth through anatomic, morphologic, and histologic descriptions. Examination of 131 skull samples, including 110 museum skull specimens and 21 harvested skulls, revealed the erupted tusks surrounded by maxillary bone over the entire length of their bone socket insertion, and are thus more accurately termed caniniform or canine teeth. The anatomy, morphology, and development of vestigial teeth in five skull samples are more fully described and documented. Vestigial tooth samples included 14 embedded pairs or individual teeth that were partially exposed or removed from the maxillary bone. Their location was posterior, ventral, and lateral to the tusks, although male vestigial teeth often exfoliate in the mouth lodging between the palatal tissue and underlying maxillary bone. Their myriad morphologies, sizes, and eruption patterns suggest that these teeth are no longer guided by function but rather by random germ cell differentiation and may eventually cease expression entirely. The conclusions reached are that the narwhal tusks are the expression of canine teeth and that vestigial teeth have no apparent functional characteristics and are following a pattern consistent with evolutionary obsolescence. Anat Rec, 2012. \u00a9 2012 Wiley Periodicals, Inc.<\/jats:p>","DOI":"10.1002\/ar.22449","type":"journal-article","created":{"date-parts":[[2012,4,3]],"date-time":"2012-04-03T09:11:25Z","timestamp":1333444285000},"page":"1006-1016","source":"Crossref","is-referenced-by-count":46,"title":["Vestigial Tooth Anatomy and Tusk Nomenclature for <i>Monodon Monoceros<\/i>"],"prefix":"10.1002","volume":"295","author":[{"given":"Martin T.","family":"Nweeia","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Frederick C.","family":"Eichmiller","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter V.","family":"Hauschka","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ethan","family":"Tyler","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James G.","family":"Mead","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Charles W.","family":"Potter","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David P.","family":"Angnatsiak","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pierre R.","family":"Richard","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jack R.","family":"Orr","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sandie R.","family":"Black","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2012,3,30]]},"reference":[{"key":"e_1_2_6_2_1","doi-asserted-by":"publisher","DOI":"10.1671\/2453-25"},{"key":"e_1_2_6_3_1","doi-asserted-by":"publisher","DOI":"10.1016\/0041-0101(67)90157-2"},{"key":"e_1_2_6_4_1","first-page":"162","article-title":"Asymmetry in Cetaceans","volume":"8","author":"Arvy L","year":"1977","journal-title":"Invest Cetacea"},{"key":"e_1_2_6_5_1","first-page":"42","article-title":"Le Narval* (Monodon monoceros L.)","volume":"73","author":"Arvy L","year":"1978","journal-title":"Acta Zool Pathol Ant"},{"key":"e_1_2_6_6_1","first-page":"52","volume-title":"Oral development and histology","author":"Avery J","year":"2002"},{"key":"e_1_2_6_7_1","first-page":"55","volume-title":"Illustrated dental embryology, histology, and anatomy","author":"Bath\u2010Balogh M","year":"1997"},{"key":"e_1_2_6_8_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1525-142X.2002.02033.x"},{"key":"e_1_2_6_9_1","first-page":"287","volume-title":"Oral anatomy, embryology and histology","author":"Berkovitz BKB","year":"2002"},{"key":"e_1_2_6_10_1","doi-asserted-by":"publisher","DOI":"10.1139\/z81-319"},{"key":"e_1_2_6_11_1","first-page":"240","volume-title":"Orban's histology and embryology","author":"Bhaskar SN","year":"1990"},{"key":"e_1_2_6_12_1","first-page":"139","article-title":"Rows of small teeth in ziphioid whales","volume":"31","author":"Boschma H","year":"1951","journal-title":"Zool Meded"},{"key":"e_1_2_6_13_1","doi-asserted-by":"publisher","DOI":"10.1016\/0003-9969(90)90027-8"},{"key":"e_1_2_6_14_1","first-page":"552","article-title":"The cetacean of the Greenland Seas","volume":"35","author":"Brown R","year":"1868","journal-title":"Proc Zool Soc London"},{"key":"e_1_2_6_15_1","first-page":"350","volume-title":"Notes and jottings from animal life","author":"Buckland FT","year":"1882"},{"key":"e_1_2_6_16_1","doi-asserted-by":"publisher","DOI":"10.1515\/mamm.1968.32.2.192"},{"key":"e_1_2_6_17_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7998.1985.tb03529.x"},{"key":"e_1_2_6_18_1","first-page":"32","article-title":"Upper canine teeth of muntiacus with particular reference to M. Reevesi","volume":"62","author":"Chapman NG","year":"1998","journal-title":"Z Saugetierkd"},{"key":"e_1_2_6_19_1","first-page":"41","article-title":"On the Narwhal\u2014On the Skeleton of the narwhal","author":"Clark JW","year":"1871","journal-title":"Proc Zool Soc London"},{"key":"e_1_2_6_20_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-3642.1985.tb01500.x"},{"key":"e_1_2_6_21_1","unstructured":"1825 De L'Institut National De France Paris MF Cuvier Des dents des mammiferes: considerees comme caracteres zoologiques 245"},{"key":"e_1_2_6_22_1","first-page":"230","article-title":"De L'Histoire naturelle des cetaces","volume":"10","author":"Cuvier MF","year":"1836","journal-title":"Libraire Encyclopedique de Roret"},{"key":"e_1_2_6_23_1","doi-asserted-by":"publisher","DOI":"10.1671\/0272-4634(2002)022[0122:CADAOA]2.0.CO;2"},{"key":"e_1_2_6_24_1","first-page":"77","article-title":"Two new species of fossil walruses from the upper Pliocene San Diego Formation, CA. Proc","volume":"29","author":"Demere TA","year":"1994","journal-title":"San Diego Soc Nat His"},{"key":"e_1_2_6_25_1","doi-asserted-by":"publisher","DOI":"10.1016\/0030-4220(77)90255-9"},{"key":"e_1_2_6_26_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.1950.0013"},{"key":"e_1_2_6_27_1","doi-asserted-by":"publisher","DOI":"10.1007\/s002270050161"},{"key":"e_1_2_6_28_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1386-1425(97)00180-7"},{"key":"e_1_2_6_29_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ea.22.050194.002223"},{"key":"e_1_2_6_30_1","first-page":"153","article-title":"Vestigial teeth in specimens of Cuvier's Whale (Ziphius cavirostris) stranded on the Scottish coast","volume":"222","author":"Fraser FC","year":"1936","journal-title":"Scottish Naturalist"},{"key":"e_1_2_6_31_1","first-page":"498","article-title":"Remarques sur la Dentition du Narval","volume":"2","author":"Gervais P","year":"1873","journal-title":"J Zool"},{"key":"e_1_2_6_32_1","first-page":"1","article-title":"Notes on a voyage to the Greenland Sea in 1888","volume":"13","author":"Gray R","year":"1889","journal-title":"Zoologist"},{"key":"e_1_2_6_33_1","first-page":"86","volume-title":"Encyclopedia of evolution","author":"Hall BK","year":"2002"},{"key":"e_1_2_6_34_1","first-page":"249","article-title":"A postglacial walrus from Bathurst Island, Northern Territories","volume":"80","author":"Harington CR","year":"1875","journal-title":"Can Field Nat"},{"key":"e_1_2_6_35_1","first-page":"36","volume-title":"Marine mammals","author":"Harrison RJ","year":"1965"},{"key":"e_1_2_6_36_1","first-page":"145","volume":"4","author":"Hay KA","year":"1989","journal-title":"Narwhal\u2014Monodon monoceros Linnaeus, 1758."},{"key":"e_1_2_6_37_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstl.1813.0021"},{"key":"e_1_2_6_38_1","first-page":"301","article-title":"On the age characteristics and anatomy of the tusk dugong dugon","volume":"30","author":"Kasuya T","year":"1978","journal-title":"Sci Rep Whales Res Inst"},{"key":"e_1_2_6_39_1","first-page":"188","volume-title":"Embryology of the mouth and related structures","author":"Kitamura H","year":"1989"},{"key":"e_1_2_6_40_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0142-9612(98)00195-1"},{"key":"e_1_2_6_41_1","first-page":"114","article-title":"Pliocene tusked odobenids (Mammalia: Carnivora) in the Western North Pacific and their paleobiogeography","volume":"21","author":"Kohno N","year":"1995","journal-title":"Bull Natl Sci Mus"},{"key":"e_1_2_6_42_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0016-6995(91)80010-W"},{"key":"e_1_2_6_43_1","doi-asserted-by":"publisher","DOI":"10.1017\/S0022336000019855"},{"key":"e_1_2_6_44_1","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.title.17555"},{"key":"e_1_2_6_45_1","doi-asserted-by":"publisher","DOI":"10.1002\/ar.20539"},{"key":"e_1_2_6_46_1","volume-title":"Permar's oral embryology and microscopic anatomy","author":"Melfi RC","year":"1988"},{"key":"e_1_2_6_47_1","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9780511565298"},{"key":"e_1_2_6_48_1","first-page":"19","volume-title":"Human oral embryology and histology","author":"Mjor IA","year":"1986"},{"key":"e_1_2_6_49_1","first-page":"155","volume-title":"Dental and oral tissues, an introduction","author":"Moss\u2010Salentijn L","year":"1990"},{"key":"e_1_2_6_50_1","first-page":"224","article-title":"Odobenocetops peruvianus, the walrus\u2010convergent Delphonoid (mammalia: cetacea) from the early Pliocene of Peru","volume":"93","author":"Muizon C","year":"2002","journal-title":"Smithson Contrib Paleobiol"},{"key":"e_1_2_6_51_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1096-3642.2002.00015.x"},{"key":"e_1_2_6_52_1","doi-asserted-by":"crossref","unstructured":"MullerGB WagnerGP.1991.Novelty in evolution: restructuring the concept. In: Annual review of ecology and sytematics22:229\u2013256.","DOI":"10.1146\/annurev.ecolsys.22.1.229"},{"key":"e_1_2_6_53_1","first-page":"88","article-title":"On pearls in elephant tusks","author":"Nagendren A","year":"1970","journal-title":"J Wildl Prot Soc Ceylon"},{"key":"e_1_2_6_54_1","first-page":"38","volume-title":"Ten Cate's oral histology: development, structure, and function","author":"Nanci A","year":"2003"},{"key":"e_1_2_6_55_1","volume-title":"Evolutionary innovations","author":"Nitecki MH","year":"1990"},{"key":"e_1_2_6_56_1","doi-asserted-by":"publisher","DOI":"10.5479\/si.097884601X.16"},{"key":"e_1_2_6_57_1","first-page":"20","volume-title":"Human tooth and dental arch development","author":"Ooe T","year":"1981"},{"key":"e_1_2_6_58_1","doi-asserted-by":"publisher","DOI":"10.1038\/049257a0"},{"key":"e_1_2_6_59_1","first-page":"412","article-title":"Mammals and birds along the east coast of Greenland. Fisheries Research Board of Canada","volume":"1206","author":"Pedersen A","year":"1931","journal-title":"Translation"},{"key":"e_1_2_6_60_1","first-page":"349","article-title":"The narwhal","volume":"2","author":"Pennent T","year":"1792","journal-title":"Artic Zool"},{"key":"e_1_2_6_61_1","first-page":"28","volume-title":"Permar's oral embryology and microscopic anatomy","author":"Permar D","year":"1988"},{"key":"e_1_2_6_62_1","first-page":"153","article-title":"Some comments on the tusk and the shape of the flukes in the narwhal (Monodon monoceros)","volume":"15","author":"Pilleri G","year":"1983","journal-title":"Invest Cetacea"},{"key":"e_1_2_6_63_1","first-page":"88","volume-title":"Fundamentals of oral histology and embryology","author":"Provenza V","year":"1988"},{"key":"e_1_2_6_64_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0003-9969(98)00077-6"},{"key":"e_1_2_6_65_1","doi-asserted-by":"publisher","DOI":"10.4102\/koedoe.v42i2.232"},{"key":"e_1_2_6_66_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0003-9969(00)00068-6"},{"key":"e_1_2_6_67_1","doi-asserted-by":"publisher","DOI":"10.2307\/3503952"},{"key":"e_1_2_6_68_1","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.title.49370"},{"key":"e_1_2_6_69_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0169-5347(98)01491-8"},{"key":"e_1_2_6_70_1","first-page":"1","volume-title":"Social organization and behavior of the narwhal, Monodon monoceros L. in lancaster sound, pond inlet, and trembaly sound, northwest territories","author":"Silverman HB","year":"1979"},{"key":"e_1_2_6_71_1","first-page":"1","article-title":"A hypothesis on the homology of proboscidean tusks based on paleontological data","volume":"2895","author":"Tassy P","year":"1987","journal-title":"Am Mus Novit"},{"key":"e_1_2_6_72_1","first-page":"7","article-title":"Growth increments in teeth of diictodon","volume":"34","author":"Thackery JF","year":"1991","journal-title":"Koedoe"},{"key":"e_1_2_6_73_1","first-page":"55A","article-title":"Whales with hind limbs and sea cows with tusks from the eocene of Pakistan","volume":"12","author":"Thewissen JG","year":"1992","journal-title":"J Vertebr Paleontol"},{"key":"e_1_2_6_74_1","first-page":"696","volume-title":"Mammals of the U.S.S.R. and adjacent countries","author":"Tomlin AG","year":"1957"},{"key":"e_1_2_6_75_1","doi-asserted-by":"publisher","DOI":"10.1006\/jasc.2002.0852"},{"key":"e_1_2_6_76_1","first-page":"316","volume-title":"Encyclopedia of marine mammals","author":"Uhen MD","year":"2008"},{"key":"e_1_2_6_77_1","doi-asserted-by":"crossref","DOI":"10.5962\/bhl.title.50302","volume-title":"Ost\u00e9ographie des c\u00e9tac\u00e9s vivants et fossiles comprenant la description et l'iconographie du squelette et du syst\u00e8me dentaire de ces animaux, ainsi que des documents relatifs \u00e0 leur histoire naturelle","author":"Van Beneden PJ","year":"1880"},{"key":"e_1_2_6_78_1","doi-asserted-by":"publisher","DOI":"10.1671\/0272-4634(2007)27[521:LCSGFI]2.0.CO;2"}],"container-title":["The Anatomical Record"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.wiley.com\/onlinelibrary\/tdm\/v1\/articles\/10.1002%2Far.22449","content-type":"unspecified","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/ar.22449","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,8]],"date-time":"2025-10-08T19:06:48Z","timestamp":1759950408000},"score":1,"resource":{"primary":{"URL":"https:\/\/anatomypubs.onlinelibrary.wiley.com\/doi\/10.1002\/ar.22449"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2012,3,30]]},"references-count":77,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2012,6]]}},"alternative-id":["10.1002\/ar.22449"],"URL":"https:\/\/doi.org\/10.1002\/ar.22449","archive":["Portico"],"relation":{},"ISSN":["1932-8486","1932-8494"],"issn-type":[{"value":"1932-8486","type":"print"},{"value":"1932-8494","type":"electronic"}],"subject":[],"published":{"date-parts":[[2012,3,30]]}}}